Tuesday, 28 November 2017

In a
previous post, I presented an aurochs reconstruction I did recently. I wrote
that it matches 100% what I imagine a Holocene European aurochs to have looked
like, and that there are obvious similarities to many Lidia bulls (Iberian fighting cattle). Of course there is the danger of a Pygmalion effect: I might
draw my aurochs Lidia-like, because I consider Lidia aurochs-like. This cannot
be ruled out completely. But I always try to follow the evidence and not to be guided
by preconceptions when doing illustrations. In the post linked above, I
explained why I drew the bull the way I did. Here is the illustration I am
talking about:

Actually,
the fact that Lidia is one of the least derived cattle breeds left today did
not just come to my mind because of that drawing, but was apparent to me right
from the beginning when I started to become interested in the aurochs. I use to
explain the looks of the aurochs to people as “much like a big, long-legged and
large-horned Iberian fighting bull”. Also, aurochs expert Cis van Vuure comes
to the conclusion that this Iberian breed is the most aurochs-like breed in his
2005 book “Retracing the aurochs:
History, morphology and ecology of an extinct wild ox”. And also Lutz Heck considered
the breed very reminiscent of the extinct wild form and relied heavily on it in
his breeding project (his lineage diminished, modern Heck cattle most likelyhas no influence from the Iberian fighting bull).

I am going
to sum up the aurochs-like pro’s of Lidia cattle:

- Muscular,
athletic body shape very reminiscent of wild bovines

- High
processus spinosi in the shoulder region (“hump”), probably as large as in the
aurochs

- Often
elongated skull shape, straight to convex snout and often prominent forelocks,
sometimes also on neck and face

- Horn
shape reminiscent of that of the aurochs with an inwards curve, although
usually not entirely as prominent as in the aurochs

- Short
dewlap and scrota and small udders in less derived individuals

- Robust
landrace, used to live outside with little husbandry all year round

Especially
the first four points are very advantageous for “breeding-back”, as those
traits are rarely found in any other primitive breeds to the same extent. All
of them give primitive Lidia individuals a very aurochs-like appearance, and I
am going to link some of those less-derived aurochs-like Lidia individuals
here:

For a video
of aurochs-like Lidia bulls, go here for example. Another video of bulls with
an exquisite anatomy and also respectable horns is here.

It is not
only their less-derived anatomy and looks that make Lidia special, but also
their hardiness and independency. As I reported in 2013, the three Lidia cows
in the German Lippeaue reserve were the only cattle that were never seen to
make use of supplementary feeding.

Despite of
their many advantages, this breed also has considerable disadvantages. First of
all, Lidia are quite small cattle. According to the Domestic Animal Diversity
Information System, bulls reach a withers height of 130cm, and cows 110, which
is 10-15cm smaller than Heck cattle and almost half a meter smaller than an
average Holocene European aurochs. But even more important, Lidia cattle are
being bred for aggression and “fighting spirit” and consequently are very
aggressive and nervous, difficult to handle and often attack easily. Additionally
to these two major disadvantages, Lidia bulls also tend to be short-legged
compared to aurochs.

But of
course it is the task of crossbreeding and selection to eradicate such negative
traits, just as too small horns or undesired colour schemes. This has been
tried in the Lippeaue, where three pure Lidia cows have been used for breeding.
But they were not really satisfied with the results. For once, the crossbreeds
remained comparably small, even if they had portions of Chianina or Sayaguesa, two
large breeds, in their genome. The crossbreeds also displayed overly nervous or
aggressive behaviour. This goes for the half-Lidia bulls as much as those that
were only quarter Lidia such as Latino. One half-Lidia bull was sold to
Hortobagy, Larus (which looked quite good), and Istvan Sandor it was the most
aggressive bull he ever experienced. Another half-Lidia bull was sold to
Denmark and slaughtered because he was too difficult to handle. I had a look at
many Lidia cross individuals in Matthias Scharf’s photo archive, and many of
them did look more muscular than crossbreeds without Lidia, but they also
tended to develop a rather elongated trunk and did not possess the slim
athletic waist of pure Lidia. As a side note, even pure Lidia bulls often grow rather
hefty at higher age, and the crossbreeds seemingly tended to go that direction
right from the beginning. So crossing-in Lidia was not that successful as
hoped, which is why the ABU diluted the influence of this breed in their herds,
and Matthias Scharf even said that he considered most Lidia-crosses to be
“small, ugly and mean”. Currently, they have one good half-Lidia cow in their
herd that also has a comparably relaxed behaviour.

Larus, a cross between a Lidia cow and a Dutch Heck bull (photo by Istvan Sandor)

So does
this mean that Lidia is, despite its obvious qualities, not a useful breed for
“breeding-back”? The Tauros Programme, for example, will not use this breed,
especially because of its behaviour. Considering the difficulties in handling
Lidia crosses, and the disappointing looks of many of the cross results
concerning size and body shape, I would say that Lidia is indeed perhaps not
worth the effort in conventional crossbreeding and keeping them in grazing
projects.

Lidia x (Heck x Chianina) cow

Nevertheless,
I also think that it would be a shame not to make use of the potential that is
in the breed. Maybe the crossbreeding results are too double-edged to speak of
a “successful use” of this breed in crossbreeding, but perhaps a project
working exclusively with good, useful individuals of that breed and trying to
eradicate the negative traits of Fighting cattle would be fruitful. I am
thinking of a project that would at first try to find a sufficient number of
aurochs-like Lidia cattle such as those on the photos I linked above that have
the right colour setting, an athletic body, elongate skulls and good horns, and
trying to acquire individuals that are as large as possible, as long-legged as
possible and perhaps not overly aggressive or nervous. For this, I would try to
search breeding sites all over Spain and Portugal and also include the Casta-Navarra
breed. Selecting them for couple of generations for the good traits and against
the three undesirable Lidia traits (small size, short-leggedness,
aggression/nervousness) might produce some good results. When selecting against
their aggressive and nervous behaviour, I would try to pay attention to not
domesticate them any further by selecting explicitly on tame and docile
behaviour, as pleiotropic effects can affect other traits as well as we see in
all domestic animals (see the Dedomestication series). The question is, then,
how to handle the cattle. It would be very difficult to next to impossible to
keep them like in grazing projects or private farms. It might be wisest to
handle them just like other Lidia breeders do, or perhaps like American farmers
keep their bison, also with the necessary equipment. After all, a true wild
aurochs probably could not be handled much differently, and it is no
coincidence either that most natural grazing projects work with cattle instead
of wisent. Of course it would be more advantageous if their behaviour was
suitable for grazing projects, as grazing projects provide the most abundant
opportunities to spread the breed and keep the animals under semi-natural
conditions in herds of sufficient size, but this post focuses only on a scheme
to breed a very aurochs-like strain of Lidia and not how to spread it. If it is
indeed possible to breed a more relaxed and less dangerous behaviour but
maintaining the wild cattle-like body conformation, it would be pleasant of
course and perhaps enable the use of these cattle in grazing projects. However,
I think it is questionable if this is possible within a few generations as
fighting cattle have been bred for aggression for dozens of generations.

The idea of
establishing an aurochs-like cattle population using Iberian Fighting cattle
only is not new. Lutz Heck considered this breed almost indistinguishable from
the aurochs, so that it is surprising that he still bred with other breeds as
well. Cis van Vuure also proposed a project working with the most primitive
Lidia cattle only in order to achieve maximum resemblance to the aurochs. I
also remember that in email conversations it was once considered to put such a
project into practise, but as several years passed now and I discovered no
mention of it on the web I assumed this idea has died a silent death.

Also, I
have my doubt that it is possible to achieve all aurochs-like traits using
Lidia only. While some Lidia grow larger than the norm, I would be very surprised
if selective breeding is able to raise the size range from 130cm for bulls to
160cm at least within a couple of generations. The same goes for horn size –
while Lidia are not small-horned, the largest horns of this breed are still
only at the lower size range of aurochs horns, perhaps still a bit too small
when considering the size of the horn sheath. Also, the breed might need a
boost in leg length, even after selective breeding. Furthermore, the problem
with breeding for quantitative traits is that it takes comparably long and other
traits might get neglected, including also genetic diversity (Lidia is already
inbred).

Thus, crossing-in
breeds that might help to overcome these deficiencies and also to increase
genetic diversity is suggestive. The first option that comes to my mind are
Chianina or Maltese cattle that would add the large size, slender posture and
long legs. Maltese cattle have the advantage of introducing colour dilution
alleles to a lesser extent and having longer snouts, but are far less
available, with not even hundred individuals and only on Malta. The big
disadvantage both breeds have are the very small horns. The horn size of Lidia
already has to be improved (although some individuals, like these young bulls,
have comparably large horns), and crossing-in one of these breeds would
introduce alleles for very small horns, and since horn size is probably
controlled by several gene loci it would be even more difficult to breed for
the desired horn size. So it would be advantageous to add another breed to increase
horn volume. Which breed would be best to do this job? One idea would be very
large-horned Texas Longhorn with the right colour setting. However, the
disadvantage would be that they are small-sized as well and have very
outwards-facing horns. Another possibility are large-horned Heck cattle from
lineages with a comparably stable inheritance; their disadvantages are their very
domestic, heavy body and paedomorphic skulls. Using Watussi would greatly
increase horn size, but also introduce many undesired traits concerning body
shape, dewlap, colour, fleshy hump, skull shape et cetera, which is why this
breed would have to be used wisely and with patience. If one cannot decide
among existing stable breeds which one to chose in order to increase horn size,
one could wait for other projects that are already in progress now to produce
results that can be used for this job. Such as the Auerrind project, which has
just started and is using Watussi. Perhaps in 15 years, when also a Lidia
project would see first results, there might already be good Watussi-influenced
Auerrind cattle that could be used in order to increase horn size in an
improved Lidia strain without introducing too many undesired traits. This is
just brainstorming – in practice it depends on many factors of course.

I would not
suggest to start crossbreeding right from the beginning and to a larger extent,
otherwise I would not be talking about a Lidia project here. It is about
preserving, concentrating and improving the aurochs-like traits present within
the Lidia population, and using other breeds only after a phase of selection
when selective breeding has been proven to not being able to reach body size,
leg length and horn size to a desirable degree. Another positive side effect of
crossing-in other breeds would be that at least some individuals might display
a more relaxed behaviour. I would use other breeds only carefully and perhaps
use only F1 crossbred individuals in order to not diminish the advantageous
Lidia traits. The point at which I would start to make use of such crossbreeds
is when selective breeding has been shown to not reach all traits to a
desirable extent, which would be after a couple of generations and therefore
about 15 years or so (which would equal about five generations).

How could
such an aurochs-like Lidia strain be named? For Maremmana, some people like to
speak of the optically less derived individuals or herds of “Maremmana
primitiva”, although those Maremmana primitiva do not represent one gene pool
or breeding line. In the case of this Lidia lineage, on the other side, it
would be one population/lineage. So perhaps calling them “Lidia primitiva”
would be an idea.

I think
that such a Lidia project could achieve a high level of authenticity regarding
looks and anatomy and perhaps also behaviour (yes, exaggerated aggression is
not “natural” behaviour; but tameness and docility are not either; you
certainly could not have used an aurochs for draft work or have milked them).
There is also the chance that with selection against exaggerated aggression and
crossing-in of other breeds Lidia primitiva might once have a behaviour that is
suitable to be used in grazing projects and other aurochs projects.

This post
is just a thought-experiment, but I hope that it inspires and perhaps other
people have similar ideas in order to make use of the potential we find in
Spanish fighting cattle.

Saturday, 18 November 2017

This title
is surely a bit provocative – it is of course zoological consensus that the Asiatic
Przewalski’s horse, Equus ferus
przewalskii, is the last living genuine wild horse that is extant today
after the last western wild horses disappeared. However, advocates of a number
of horse breeds that they purport as living remnants of the European wild horse
or at least being strongly influenced by original wild horses, sometimes put
the status of the Przewalski’s horse as a genuine wild animal into question.
They argue that decades of breeding in captivity and introgression from
domestic horses has altered the nature of the Przewalski’s horse, and claim
that the situation is comparable to what has happened to their favoured “wild”
breed: an original wild population has been influenced by domestic horses and artificial
selection. As a consequence, they argue that if the Przewalski’s horse still
deserves status as a genuine wild horse, which is zoological consensus, then so
does their breed of choice. However, the often purported background stories for
those breeds being “near-wild horses” are not tenable after objective
examination for a number of reasons (see here), but what about the arguments
against the original, wild status of the Przewalski’s horse?

The lineage
of the Przewalski’s horse separated from that of domestic horses several
millennia ago. The exact point of separation varies from study to study,
depending on the molecular chronometer and its calibration. The maximum I found
was 160.000 [1] and the minimum 38.000 [2] years ago. This comparably long
reproductive separation resulted in a different karyotype, the Przewalski’s
horse having 23 pairs of chromosomes and the domestic horse 24 due to a fission
or fusion (depending on what is the plesiomorphic state), but they intermix
readily and without fertility problems. In the millennia of living side by side
in the Eurasian steppe, the Przewalski’s horse contributed genetically to the
domestic stock (which is not only genetically [3] but also optically apparent,
f.e. see some Mongolian horses), and vice versa. The Przewalski’s horse gene
pool was introgressed by domestic horses, especially in the 20th
century. Photographs of wild herds from 1954 showed individuals of divergent
colours (Wikipedia), indicating admixture. The whole modern population descends
from 13 founding individuals, one of them was a domestic Mongolian stallion
[4]. Does this mean that the original, genuine Przewalski’s horse is lost and
the modern population is an altered result of intermixture?

Orlando et
al. 2015 made a genomic study compromised of a large sample of Przewalski’s
horses, post and prior to the bottleneck (including the holotype specimen),
domestic horses plus a late Pleistocene wild horse as outgroup. The result is
that although there are genetic traces of intermixture, also including such
having an effect on the phenotype such as an allele associated with increased
withers height, there are still lineages in the population that are virtually
free of admixture[3]. Also, height is a highly multifactorial trait, therefore
it cannot be claimed that Przewalski’s horses are taller now due to admixture
because of one allele – the average withers height is still between 122-142cm
according to English Wikipedia, between 120-146 according to German Wikipedia
(note that there is also sexual dimorphism in size). Przewalski’s horses are
still uniform in their typical colour, sturdy build, robust head shape, erect
and short mane, short-haired tail basis, a very lightly coloured almost white
winter coat, and other typical morphologic differences to domestic horses such
as thicker hooves (Wikipedia). I would even say that domestic cattle left a
bigger trace on modern American bison than domestic horses did on the
Przewalski’s horse, yet nobody is questioning the bison’s status as a wild
animal. Also I found no source stating that Przewalski’s horses with a domestic
karyotype have been observed.

Yes, the
Przewalski’s horse seemingly intermixed with domestic horses continuously after
their point of separation, but I see no compelling evidence that this fact
altered the genetic integrity of this wild subspecies. Furthermore, domestic
animals introgressed the gene pool of their wild counterparts everywhere they
shared the habitat – this evident in European wild boar that show deviant,
domestic colours and there is also the hypothesis that American wolves
inherited black and other colour variants from domestic dog introgression
several millennia ago (this might also explain blue-eyed wolves). Yet nobody is
calling their wild animal status into question.

If
Przewalski’s horses indeed lost part of their wild animal nature due to
domestic introgression and being bred in captivity for a number of decades, it
might be helpful to look at a checklist of aspects typical signs of
domestication:

- Earlier
maturity and increased litter size (the latter aspect is not true of domestic
horses either, so let us ignore it for now)

Przewalski’s
horses do not show any signs of morphological paedomorphy, not even if you
compare photos of the early 1900s to modern individuals. Przewalski’s horses
still always have the robust, donkey-like skull with small eyes and their
proportions do not seem to be altered as well. I have not found any remarks in
the literature stating that Przewalski’s horses as a whole lost brain volume;
domestic horses have about 14% less brain volume than Przewalski’s horses [5]. Captive
Przewalski’s horses also have 14% less brain capacity than wild counterparts
[5]. Since there are no separate genetic lineage between wild and captive
Przewalski’s horses, this should be applicable to phenotypic plasticity.
According to Wikipedia, earlier maturity in captive Przewalski’s horses has
been reported, but explained with better nutritional conditions in zoos than in
the wilderness and as far as I know the same phenomenon can be observed in
other zoo animals. The behaviour of Przewalski’s horses and domestic horses is
well comparable, but Przewalski’s horses have a way higher aggression potential
than domestic horses, especially the stallions. This is universal for this
subspecies and evident in zoos as well as grazing projects. I once was told
that zookeepers are more afraid of Przewalski’s horses than lions. Przewalski’s
horses can be tamed and ridden to a certain degree, but this is also true for
zebras, including the quagga.

There are
not any novel traits found in any Przewalski’s horse, such as a new colour
variant, or fur modifications. There are occasionally individuals showing a
white streak along the face or white socks, which is applicable to
introgression from domestic horses.

Looking at
some deer populations which have been kept in game parks for many generations,
we see incipient signs of domestications, such as new colour variants or
typical domestic spotted patterns, or beginning paedomorphic skull shapes – you
can find this in some roe deer, red deer and fallow deer in European game parks
and this is what I would call an early state of slow domestication. But we do
not see that at all in Przewalski’s horses.

All in all
I think there is not one compelling reason to claim that the original
Przewalski’s horse is gone, that it has been altered by man and hybridization,
or that it is on the edge of domestication. I see nothing that calls their
status as a genuine wild animal seriously into question, especially when we
look at other wild animals. And even if the critics were right, it would not
make any of the domestic horse breeds praised as near-wild horses “wilder” than
they are (or not are, actually).

About this blog

This blog is on everything related to the so-called “breeding-back” of extinct animals: From the extinct animals themselves, over their often domestic descendants and dedomestication to news and facts about various breeding-back projects, reports and photos from my own breeding-back related trips. I try to have a balanced and fact-based approach to this subject and to dismantle many of the popular myths. Enjoy!

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About me

My major interest always have been extinct animals, from dinosaurs to Pleistocene megafauna and more recent extinctions. Besides that I am interested in evolution, genetics and ecology.
I am also an amateur animal artist, making drawings and models mostly of extinct animals.